Germination Ecophysiology of the Mesic Deciduous Forest Herb Polemonium reptans var. reptans (Polemoniaceae)

Abstract Seeds of Polemonium reptans var. reptans , a perennial herb of mesic deciduous forests in eastern North America, mature in late May-early June, and a high percentage of them are dormant. Seeds afterripened (came out of dormancy) during summer when kept in a nylon bag under leaves in a nonheated greenhouse or on wet soil in a 30/15°C incubator. The optimum temperature for germination of nondormant seeds was a simulated October (20/10°C) regime. In germination phenology studies in the nonheated greenhouse, 20–30% of the seeds that eventually germinated did so in October, and the remainder germinated the following February and March. Since low (5°C) winter temperatures promote some afterripening (ca. 50%) and do not cause nondormant seeds to re-enter dormancy, seeds that fail to germinate in autumn may germinate in spring. Thus, the taxon has very little potential to form a persistent seed bank. The large spatulate embryos and ability of seeds to afterripen at high temperatures means that seeds of P. reptans var. reptans have nondeep physiological dormancy, unlike many herbaceous woodland species, which have morphophysiological dormancy.     

Germination ecology of the polycarpic grassland perennials Primula veris and Trollius europaeus

The seed germination behaviour of Primula veris and Trollius europaeus , both perennial, polycarpic grassland plants was compared The species have similar-sized seeds that are dormant at dispersal Seeds buried in soil and exhumed at regular intervals showed that for both species, primary seed dormancy was overcome by cold-stratification Hence, their germination in the field should occur in spring, following dispersal, or later Seeds of P veris became dormant again in the late spring/early summer, and dormancy was broken again in the second winter Seeds of T europaeus did not exhibit such changes in dormancy
Seeds of P veris did not germinate in darkness This suggests that P veris can accumulate a persistent seed bank because buried seeds are prevented from germinating Trollius europaeus , on the other hand, germinated equally well in darkness and in light which suggests that seeds might germinate even when they are too deep in the soil for seedlings to emerge Two lines of evidence confirm this difference in seed bank behaviour (1) Primula veris was detected in the persistent seed bank of a grassland site, whereas T europaeus was not (n) After 16 months burial, 85% of the P veris seeds but only 8% of the T europaeus seeds remained viable     

Seed Germination Ecology of the Cold Desert Annual Isatis violascens (Brassicaceae): Two Levels of Physiological Dormancy and Role of the Pericarp

The occurrence of various species of Brassicaceae with indehiscent fruits in the cold deserts of NW China suggests that there are adaptive advantages of this trait. We hypothesized that the pericarp of the single-seeded silicles of Isatis violascens restricts embryo expansion and thus prevents germination for 1 or more years. Thus, our aim was to investigate the role of the pericarp in seed dormancy and germination of this species. The effects of afterripening, treatment with gibberellic acid (GA₃) and cold stratification on seed dormancy-break were tested using intact silicles and isolated seeds, and germination phenology was monitored in an experimental garden. The pericarp has a role in mechanically inhibiting germination of fresh seeds and promotes germination of nondormant seeds, but it does not facilitate formation of a persistent seed bank. Seeds in silicles in watered soil began to germinate earlier in autumn and germinated to higher percentages than isolated seeds. Sixty-two percent of seeds in the buried silicles germinated by the end of the first spring, and only 3% remained nongerminated and viable. Twenty to twenty-five percent of the seeds have nondeep physiological dormancy (PD) and 75–80% intermediate PD. Seeds with nondeep PD afterripen in summer and germinate inside the silicles in autumn if the soil is moist. Afterripening during summer significantly decreased the amount of cold stratification required to break intermediate PD. The presence of both nondeep and intermediate PD in the seed cohort may be a bet-hedging strategy.     

DORMANCY AND GERMINATION IN SEEDS OF COMMON RAGWEED WITH REFERENCE TO BEAL'S BURIED SEED EXPERIMENT

Common ragweed (Ambrosia artemisiifolia L.) was one of 19 herbaceous weedy species used by Beal in his buried viable seed experiment started in 1879. No seeds germinated during the first 35 years of the experiment when germination tests were performed in late spring, summer or early autumn. Germination did occur in seeds buried for 40 years when seeds were exhumed and tested for germination in early spring. Data obtained in more recent research provide the probable explanation for these results. Seeds of common ragweed that do not germinate in spring enter secondary dormancy by mid to late spring and will not germinate until dormancy is broken the following late autumn and winter. Thus, during the first 35 years of the experiment seeds were dormant when tested for germination, whereas seeds buried for 40 years were nondormant. Seeds buried 50 years or longer did not germinate when tested in spring, probably because they had lost viability and/or seeds germinated during burial and seedlings died.     

Dormancy-breaking and germination requirements for seeds of Symphoricarpos orbiculatus (Caprifoliaceae)

Fruits (drupes) of Symphoricarpos orbiculatus ripen in autumn and are dispersed from autumn to spring. Seeds (true seed plus fibrous endocarp) are dormant at maturity, and they have a small, linear embryo that is underdeveloped. In contrast to previous reports, the endocarp and seed coat of S. orbiculatus are permeable to water; thus, seeds do not have physical dormancy. No fresh seeds germinated during 2 wk of incubation over a 15°/6°-35°/20°C range of thermoperiods in light (14-h photoperiod); gibberellic acid and warm or cold stratification alone did not overcome dormancy. One hundred percent of the seeds incubated in a simulated summer → autumn → winter → spring sequence of temperature regimes germinated, whereas none of those subjected to a winter → spring sequence did so. That is, cold stratification is effective in breaking dormancy only after seeds first are exposed to a period of warm temperatures. Likewise, embryos grew at cold temperatures only after seeds were exposed to warm temperatures. Thus, the seeds of S. orbiculatus have nondeep complex morphophysiological dormancy. As a result of dispersal phenology and dormancy-breaking requirements, in nature most seeds that germinate do so the second spring following maturity; a low to moderate percentage of the seeds may germinate the third spring. Seeds can germinate to high percentages under Quercus leaf litter and while buried in soil; they have little or no potential to form a long-lived soil seed bank.     

A combined physical and physiological dormancy controls seasonal seedling emergence of Geranium robertianum

Temperate forest herbs with seeds exhibiting both a physical and a physiological dormancy mechanism are rare, and knowledge on the factors regulating germination of these species is fragmentary. The biennial Geranium robertianum L. grows mainly in temperate woodlands, but can also be found in exposed habitats. Seedlings of G. robertianum are known to emerge from spring until autumn, but little is known about the environmental factors regulating germination. In this study, phenology of seedling emergence and of physical dormancy loss was examined for seeds buried at shaded or sunny exposed locations. The role of temperature in regulating dormancy and germination was analysed by incubating seeds in temperature sequences simulating temperatures that seeds experience in nature. The results indicate that most seeds of G. robertianum buried in sunny conditions germinate immediately after physical dormancy loss in summer. Seeds buried in shaded conditions also lose physical dormancy mainly during summer, but remain physiologically dormant and do not germinate until late winter or early spring. Besides physical dormancy, seeds of G. robertianum also initially have a high level of physiological dormancy, which is reduced during dry storage. Physiological dormancy is reduced through chilling in winter, thus enabling the seeds to germinate at low temperatures. We conclude that a complex combination of physical and physiological dormancy ensures that G. robertianum seeds germinate in summer at exposed sites and in early spring at shaded sites.     

GERMINATION ECOLOGY OF SEDUM PULCHELLUM MICHX. (CRASSULACEAE)

Factors controlling the timing of seed germination were investigated in the small succulent winter annual Sedum pulchellum Michx. (Crassulaceae) in its natural habitat on unshaded limestone outcrops in northcentral Kentucky. At maturity in early July the dormant seeds are not dispersed but are retained in the fruits on the standing dead plants until September and October. Many, but not all, of the seeds afterripen in the fruits during summer, and at the time of dispersal some of them are dormant and some are nondormant. Germination and annual population establishment occur in September and October from seed reserves that have been in the soil for one or more years and from seeds produced in the current year. Germination of nondormant seeds may be prevented in autumn by lack of the appropriate combination of environmental factors including light, temperature and soil moisture in the seed's microsite. The effect of low winter temperatures on ungerminated seeds in the population is to induce nondormant seeds into secondary dormancy and to prevent afterripening of dormant seeds. Thus, in spring all the seeds in the population's seed reserve are dormant. During spring and summer some of these seeds afterripen, and they germinate in autumn when, and if, germination requirements are fulfilled.     

PHYSIOLOGICAL ECOLOGY OF GERMINATION OF VIOLA RAFINESQUII

Seeds of the winter annual Viola rafinesquii Greene exhibit true dormancy at the time of maturity and dispersal in mid to late spring. During the summer rest period the seeds pass from a state of true dormancy to one of relative dormancy and finally to what may be called a state of complete nondormancy. As the seeds enter relative dormancy they will germinate mostly at relatively low temperatures (10, 15, 15/6, and 20/10 C), but as after-ripening continues they gain the ability also to germinate at higher temperatures (20, 25, and 30/15 C). During June, July, and August seeds will not germinate at field temperatures even if kept continuously moist. But by September and October seeds may germinate to high percentages over a wide range of temperatures, including September and October field temperatures. This pattern of germination responses, involving breaking of true dormancy and widening of the temperature range for germination during relative dormancy, appears to be an adaptation of the species to a hot, dry season. Seeds of V. rafinesquii stored on continuously wet soil (field capacity) or on soil that was alternately wet and dried during the summer did not after-ripen at low temperatures (10, 15, 15/6, and 20/10 C) but did after-ripen fully at high temperatures (20, 25, 30/15, and 35/20 C). Thus, the high temperatures that V. rafinesquii “avoids” by passing the summer in the dormant seed stage actually are required to break seed dormancy and, therefore, are essential for completion of its life cycle.     

Seed development and germination ecophysiology of the invasive tree <Emphasis Type="Italic">Prunus serotina</Emphasis> (Rosaceae) in a temperate forest in Western Europe

Seed development, dormancy and germination of the American invasive tree species, Prunus serotina, are described for plants growing in a large forest in Belgium. Seeds of P. serotina were collected following anthesis in the first week of July and thereafter at fortnightly intervals. Seed dormancy, temperature requirements for germination and the soil seed bank were investigated. At maturation (about 105 days after anthesis), seed moisture content had decreased to around 13.7%, and 44% of the seeds had attained the capacity to germinate. Mature seeds of P. serotina exhibited physiological dormancy, germinating only after a long cold, moist stratification period. Highest germination percentage occurred in seeds treated with gibberellic acid (GA₃), at 10°C. We found no evidence that P. serotina forms a persistent seed bank but noticed a persistent seedling bank in the field.     

Evidence for physiological seed dormancy cycling in the woody shrub Asterolasia buxifolia and its ecological significance in fire‐prone systems

• Dormancy cycling is a key mechanism that contributes to the maintenance of long‐term persistent soil seed banks, but has not been recorded in long‐lived woody shrub species from fire‐prone environments. Such species rely on seed banks and dormancy break as important processes for post‐fire recruitment and recovery.
• We used germination experiments with smoke treatments on fresh seeds and those buried for 1 year (retrieved in spring) and 1.5 years (retrieved the following late autumn) to investigate whether Asterolasia buxifolia, a shrub from fire‐prone south‐eastern Australia with physiologically dormant seeds, exhibited dormancy cycling.
• All seeds had an obligation for winter seasonal temperatures and smoke to promote germination, even after ageing in the soil. A high proportion of germination was recorded from fresh seeds. but germination after the first retrieval was significantly lower, despite high seed viability. After the second retrieval, germination returned to the initial level. This indicates a pattern of annual dormancy cycling; one of the few observations, to our knowledge, for a perennial species. Additionally, A. buxifolia’s winter temperature and smoke requirements did not change over time, highlighting the potential for seeds to remain conditionally dormant (i.e. restricted to a narrow range of germination conditions) for long periods.
• For physiologically dormant species, such as A. buxifolia, we conclude that dormancy cycling is an important driver of successful regeneration, allowing seed bank persistence, sometimes for decades, during fire‐free periods unsuitable for successful recruitment, while ensuring that a large proportion of seeds are available for recruitment when a fire occurs.

     

Annual dormancy cycles in buried seeds of shrub species: germination ecology of Sideritis serrata (Labiatae)

The germination ecology of Sideritis serrata was investigated in order to improve ex‐situ propagation techniques and management of their habitat. Specifically, we analysed: (i) influence of temperature, light conditions and seed age on germination patterns; (ii) phenology of germination; (iii) germinative response of buried seeds to seasonal temperature changes; (iv) temperature requirements for induction and breaking of secondary dormancy; (v) ability to form persistent soil seed banks; and (vi) seed bank dynamics. Freshly matured seeds showed conditional physiological dormancy, germinating at low and cool temperatures but not at high ones (28/14 and 32/18 °C). Germination ability increased with time of dry storage, suggesting the existence of non‐deep physiological dormancy. Under unheated shade‐house conditions, germination was concentrated in the first autumn. S. serrata seeds buried and exposed to natural seasonal temperature variations in the shade‐house, exhibited an annual conditional dormancy/non‐dormancy cycle, coming out of conditional dormancy in summer and re‐entering it in winter. Non‐dormant seeds were clearly induced into dormancy when stratified at 5 or 15/4 °C for 8 weeks. Dormant seeds, stratified at 28/14 or 32/18 °C for 16 weeks, became non‐dormant if they were subsequently incubated over a temperature range from 15/4 to 32/18 °C. S. serrata is able to form small persistent soil seed banks. The maximum seed life span in the soil was 4 years, decreasing with burial depth. This is the second report of an annual conditional dormancy/non‐dormancy cycle in seeds of shrub species.     

Temperature conditions control embryo growth and seed germination of Corydalis solida (L.) Clairv., a temperate forest spring geophyte

Spring is often the most suitable period for seedling establishment of temperate woodland species. Different physiological mechanisms resulting in spring emergence have evolved in seeds of such plants. The aim of this study was to determine the requirements for breaking dormancy and for seed germination of the European perennial spring geophyte Corydalis solida (Fumariaceae). Ripe seeds of C. solida contain an underdeveloped embryo, consisting of no more than a clump of cells. As a consequence, the embryo has to differentiate and grow to a critical length before germination can occur. In nature, seeds are dispersed in spring, while growth of the embryo starts in the autumn and continues in winter. Germination starts in late winter, immediately after embryo growth is completed, resulting in seedling emergence in the following spring. Experiments in controlled conditions showed that temperature is the main factor controlling dormancy and germination. Incubation at autumn temperatures (15/6 °C; 20/10 °C) for at least 8 weeks is required to initiate embryo growth, while a transfer to 5 °C is needed for completion of embryo growth and germination. Growth of the embryo of C. solida occurs at different temperatures over an extended period, a feature typical of temperate forest herbs. Our results indicate that the dormancy mechanism in seeds of C. solida is very similar to mechanisms in other Corydalis species studied thus far, suggesting that stasis in the dormancy trait has occurred.     

A season- and gap-detection mechanism regulates seed germination of two temperate forest pioneers

The survival of seedlings in temperate climate habitats depends on both temporal and spatial factors. The interaction between an internal seed dormancy mechanism and the ruling environmental conditions allows accurate cueing of germination. We analysed how environmental signals interact in seeds of temperate forest pioneer species, increasing the seed's chances of germinating in the right place at the right time. Digitalis purpurea and Scrophularia nodosa are two small-seeded herbaceous species that typically grow in vegetation gaps in European temperate forests. Seeds of both species are partially dormant at the time of dispersal in summer. This primary dormancy is released in autumn and early winter, resulting in a minimal level of physiological dormancy by late winter and early spring. We observed that physiological dormancy was induced again in seeds exhumed in late spring and in summer. Experiments in laboratory conditions revealed that primary dormancy in seeds of S nodosa was broken by cold stratification, whereas primary dormancy in D. purpurea seeds was broken by both a cold and a warm stratification. The two species differed in their response to the tested gap-detection signals, as light was the most important factor stimulating germination of D. purpurea, and seeds of S. nodosa germinated best when subjected to daily fluctuating temperatures. This study clearly indicates that the ability to germinate in response to gap-detection signals changes seasonally in temperate forest pioneers. Additionally, seeds of both species responded differently to these environmental signals, probably reflecting differences in the regeneration niche.     

Morphological side effects of using gibberellic acid to induce germination: consequences for the study of seed dormancy

To assess the evolutionary significance of persistent seed banks, phenotypes of naturally germinating seeds must be compared with those that remain dormant under the same environmental conditions. Dormant seeds can often be induced to germinate by application of gibberellic acid (GA). However, this method is valid only if there are no phenotypic “side effects” of GA that could confound comparisons between dormant and naturally germinating seeds. We examined this assumption in Lesquerella fendleri, a short-lived perennial mustard of the desert Southwest. We exposed 3840 seeds from 16 maternal sibships to two different GA treatments (0 or 1 g/L GA) in two different germination environments (greenhouse and growth chamber), and measured germination and postgermination traits. As expected, application of GA increased germination. GA also had strong and long-lasting effects on seedling morphology. Seeds that received GA developed into seedlings that were taller, with fewer but longer leaves, than seeds that did not receive GA. Effects of GA on both dormancy and postgermination traits varied among maternal sibships. Our results indicate that for this species and this concentration of GA, morphological effects can be substantial. Further study is required to determine whether such side effects are found for lower concentrations of GA, or under conditions that encourage faster seedling growth. Nonetheless, the present results illustrate the importance of testing potential confounding effects of GA in studies of the evolution of seed dormancy and its influence on postgermination traits.     

R.M. Cowling,D.M. Richardson & S.M. Pierce (eds.) 1977. Vegetation of Southern Africa.

We determined the role of bird dispersal in seed and seedling dynamics of the tree Kalopanax pictus from 7 years of observing seed rain and seedling emergence in a broad-leaved deciduous forest in central Japan. We also performed an experiment on the influence of seed pulp on germination of seeds of K. pictus. Seeds of this species can lie dormant for several years, and this causes rather constant yearly seedling emergence in spite of irregular seed production. The spatial distribution of the seedlings that emerged each year (maximum distance from nearest conspecific seed-bearing tree of 90 m) was wider than that of gravity-dispersed seeds (max. distance of 37 m), suggesting seed dispersal by birds in winter. Emerged seedling densities at sites over 20 m from the nearest conspecific seed-bearing tree were highest in the spring of 1991, about half a year after the largest seed fall of the observation period. However, emerged seedling densities within 20 m from seed-bearing trees were highest in 1992, 1.5 years after the largest seed fall. These field observations may be explained by the experimental results on the effects of seed pulp on germination. Intact seeds germinate slowly at low germination rates, while seeds without seed pulp germinate quickly at high germination rates. Fallen seeds with seed pulp thus appear to form a seed bank near seed sources (temporal dispersal), while seeds scattered by birds appear to increase the possibility of reaching the present safe sites in distant areas with quick germination (spatial dispersal).     

Seed Treatment Optimizes Benefits of Seed Bank Storage for Restoration‐Ready Seeds: The Feasibility of Prestorage Dormancy Alleviation for Mine‐Site Revegetation

Dormant seeds of 18 species from 9 families covering a diverse range of seed dormancy syndromes and life histories from the southwest Australian biodiversity hotspot were assessed for germinability following storage at 15–25°C for 36 months. A total of 10 species with physical dormancy (PY) and 8 with either physiological dormancy (PD) or morphophysiological dormancy (MPD) were assessed as part of the study. Prior to storage, germination from dormant seeds was 1–27%, rising to 41–100% following specific dormancy‐breaking treatments. When seed dormancy was removed prior to storage for 36 months seeds from all species were found to maintain a nondormant state and germinate to a similar level to that observed at the beginning of the experiment (44–100%). Likewise, seeds that did not receive a prestorage dormancy‐breaking treatment maintained a dormant state (0–50% germination) and subsequently responded well to a dormancy‐breaking treatment immediately prior to germination assessment (49–99%). There were minimal differences in response to dormancy‐breaking treatments before and after 36 months storage (average 4–6% difference) and in the germination responses observed between both storage environments assessed (15°C/15% eRH or 15–25°C air dried). Based on these findings, storing seeds in a nondormant state does not alter germinability and this approach provides significant benefits to current seed‐based restoration programs through reduction of double handling and improved seed use efficiency.     

Dormancy cycling and persistence of seeds in soil of a cold desert halophyte shrub

Background and Aims

Formation of seed banks and dormancy cycling are well known in annual species, but not in woody species. In this study it was hypothesized that the long-lived halophytic cold desert shrub Kalidium gracile has a seed bank and dormancy cycling, which help restrict germination to a favourable time for seedling survival.

Methods

Fresh seeds were buried in November 2009 and exhumed and tested for germination monthly from May 2010 to December 2011 over a range of temperatures and salinities. Germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were investigated in the field.

Key Results

Seeds of K. gracile had a soil seed bank of 7030 seeds m⁻² at the beginning of the growing season. About 72 % of the seeds were depleted from the soil seed bank during a growing season, and only 1·4 % of them gave rise to seedlings that germinated early enough to reach a stage of growth at which they could survive to overwinter. About 28 % of the seeds became part of a persistent soil seed bank. Buried seeds exhibited an annual non-dormancy/conditional dormancy (ND/CD) cycle, and germination varied in sensitivity to salinity during the cycle. Dormancy cycling is coordinated with seasonal environmental conditions in such a way that the seeds germinate in summer, when there is sufficient precipitation for seedling establishment.

Conclusions

Kalidium gracile has three life history traits that help ensure persistence at a site: a polycarpic perennial life cycle, a persistent seed bank and dormancy cycling. The annual ND/CD cycle in seeds of K. gracile contributes to seedling establishment of this species in the unpredictable desert environment and to maintenance of a persistent soil seed bank. This is the first report of a seed dormancy cycle in a cold desert shrub.     

Sensitivity cycling in physically dormant seeds of the Neotropical tree Senna multijuga (Fabaceae)

• Cycling of sensitivity to physical dormancy (PY) break has been documented in herbaceous species. However, it has not been reported in tree seeds, nor has the effect of seed size on sensitivity to PY‐breaking been evaluated in any species. Thus, the aims of this study were to investigate how PY is broken in seeds of the tropical legume tree Senna multijuga, if seeds exhibit sensitivity cycling and if seed size affects induction into sensitivity.
• Dormancy and germination were evaluated in intact and scarified seeds from two collections of S. multijuga. The effects of temperature, moisture and seed size on induction of sensitivity to dormancy‐breaking were assessed, and seasonal changes in germination and persistence of buried seeds were determined. Reversal of sensitivity was also investigated.
• Fresh seeds were insensitive to dormancy break at wet–high temperatures, and an increase in sensitivity occurred in buried seeds after they experienced low temperatures during winter (dry season). Temperatures ≤20 °C increased sensitivity, whereas temperatures ≥30 °C decreased it regardless of moisture conditions. Dormancy was broken in sensitive seeds by incubating them at 35 °C. Sensitivity could be reversed, and large seeds were more sensitive than small seeds to sensitivity induction.
• Seeds of S. multijuga exhibit sensitivity cycling to PY‐breaking. Seeds become sensitive during winter and can germinate with the onset of the spring–summer rainy season in Brazil. Small seeds are slower to become sensitive than large ones, and this may be a mechanism by which germination is spread over time. Sensitive seeds that fail to germinate become insensitive during exposure to drought during summer. This is the first report of sensitivity cycling in a tree species.

     

Morphological dormancy in seeds of the autumn-germinating shrub Lonicera caerulea var. emphyllocalyx (Caprifoliaceae)

To better understand the germination ecophysiology of the genus Lonicera , the dormancy class, temperature requirements for embryo growth and radicle emergence and phenology of seedling emergence were determined for Lonicera caerulea var. emphyllocalyx . At maturity, seeds have an underdeveloped embryo (approximately 28% of the length of full-grown embryos). Embryos in fresh seeds grew to full length at 15, 20, 20/10 and 25/15°C within 3 weeks, but failed to grow at ≤ 10°C and at 30°C. Radicles emerged from 86–100% of freshly matured seeds in light at 15, 20, 20/10 and 25/15°C within 28 days, but failed to emerge at 10°C. Radicles emerged equally well in a 12 h photoperiod and in continuous darkness at 25/15°C. Rapid embryo growth and germination over a range of conditions indicate that seeds of this taxon have morphological dormancy (MD); this is the first report of MD in a species of Lonicera . Seeds are dispersed in summer, at which time high temperatures promote embryo growth. Embryos grow to the critical length for germination in approximately 1 month; the peak of seedling emergence occurs in early autumn. Radicles emerged within 2 months from 98% of seeds buried at soil depths of 2 cm and 10 cm in the field in August in Sapporo, Japan; thus, seeds have no potential to form a persistent soil seed bank. However, seeds sown too late in autumn for embryos to grow remained viable and germinated the following summer when temperatures were high enough to promote embryo growth.     

Variation within species and inter-species comparison of seed dormancy and germination of four annual Lamium species

In an ecological context, knowledge of intra-species variation in dormancy and germination is necessary both for practical and theoretical reasons. We used four or five seed batches (replicates) of four closely related annuals co-occurring in arable fields in Sweden: Lamium amplexicaule, L. confertum, L. hybridum and L. purpureum. Seeds used for experiments stemmed from plants cultivated on two sites, each site harbouring one population of each species, thereby ensuring similar environmental history of seeds. Seeds were tested for germination when fresh and after three different pre-treatments (cold or warm stratification, or dry storage) for up to 24 weeks. Seeds were also sown outdoors. Despite substantial intra-species variation, there were clear differences between species. The general seed dormancy pattern, i.e. which environmental circumstances that affect dormancy, was similar for all species; dormancy reduction occurred during warm stratification or dry storage. Even though the response to warm stratification indicates a winter annual pattern, successful plants in Sweden were mostly spring emerged. Germination in autumn occurred, but plants survived winters poorly. Consequently, as cold stratification did not reduce dormancy, strong dormancy in combination with dormancy reduction during dry periods might explain spring germination. It is hypothesised that local adaptations occur through changes mainly in dormancy strength, i.e. how much effort is needed to reduce dormancy. Strong dormancy restricts the part of each seed batch that germinate during autumn, and thus reduces the risk of winter mortality, in Sweden.